Scientia Horticulturae, 20 (1983) 1--14 1

Elsevier Science Publishers B.V., Amsterdam -- Printed in The Netherlands

EFFECT OF CO2 ENRICHMENT ON PHOTOSYNTHESIS, GROWTH AND

YIELD OF TOMATO

STEIN NILSEN, KIRSTI HOVLAND, CHRISTIAN DONS and SONJA PREHN SLETTEN
The Phytotron, University of Oslo, Box 1066, Blindern, Oslo 3 (Norway)
(Accepted for publication 6 July 1982)

ABSTRACT

Nilsen, S., Hovland, K., Dons, C. and Sletten, S.P., 1983. Effect of CO~ enrichment on
photosynthesis, growth and yield of tomato. Scientia Hortic., 20: 1--14.

Net photosynthesis of tomato plants was measured as CO~ uptake in various light inten-
sities, COs concentrations, O5 concentrations and temperatures during short-term experi-
ments. Net photosynthesis increased significantly with increasing CO2 concentration at all
light intensities, even at the lowest one. The optimal temperature for photosynthesis in-
creased with COs enrichment. The changes in photosynthesis when the COJO2 ratio was
varied suggest that the effect of CO2 enrichment is a result of a reduction in photorespira-
tion.
To determine whether the increase in photosynthesis caused by CO2 enrichment would
produce a greater yield, tomato plants were cultivated from seed to harvest in a tightly-
closed greenhouse that was enriched with COs continuously during the entire growing-peri-
od. The control greenhouse was ventilated by openings in the roof and door. The tempera-
ture of the 2 greenhouses was kept the same. Plants enriched with CO~ showed a significant
increase in fresh and dry weight and yield of tomatoes. The results are discussed in relation
to earlier work in which CO2 enrichment was discontinued when there was a need for venti-
lation.

Keywords: CO2; light intensity; 02; photosynthesis; temperature; tomato.

INTRODUCTION

The enhancement of photosynthesis, growth and yield by CO2 enrichment

has been well established (Kramer, 1981). The effect of CO2 enrichment on
growth has been related to the increase in photosynthesis (Gaastra, 1966), re-
sulting from (1) a change in the internal photosynthetic capacity of the plant
(Hicklenton and Jolliffe, 1978), or (2) an increase in the activity of enzymes
regulating photosynthesis such as RuBP carboxylase, (3) a decrease in photo-
respiratory enzymes such as glycolic acid oxidase (Hicklenton and Jolliffe,
1980), or (4) a steeper gradient favouring CO2 diffusion into the mesophyll
(Gaastra, 1966). The developmental changes which plants cultivated in high
CO 2 undergo may also contribute to the CO2 enrichment effects (Hicklenton
and Jolliffe, 1980). The suggestion that CO2 affects the stomata (Heath and
Russel, 1954; Pallas, 1965) is still quite controversial.

0304-4238/83/$03.00 1983 Elsevier Science Publishers B.V.

 The chemical step in CO: reduction is catalyzed by the enzyme RuBP-car-
boxylase in the carbon reduction cycle. If CO: is a limiting factor, increasing
the CO 2 concentration should result in a higher production of organic matter.
However, the ratio of CO2 to 02 must be considered, since it will determine
the ratio of photorespiration, a process closely related to, and competitive
with, photosynthesis. RuBP produced in the carbon reduction cycle can be
carboxylated to form 2 molecules of 3-phosphoglycerate (PGA) in photosyn-
thesis, or through an oxidation process it can be split to form PGA and P-gly-
colate, which initiates photorespiration (Lorimer et al., 1973). A high CO2/O2
ratio will direct the reaction through photosynthesis, a low ratio through pho-
torespiration (Bowes et al., 1971; Bowes and Ogren, 1972; Badger and
Andrews, 1974). The effect of this balance is independent of the available en-
ergy from the photosynthetic light reaction. We therefore suggest that CO2 en-
richment depresses photorespiration and favors net photosynthesis in all light
intensities. This further implies that the optimal growing-conditions in CO2-
enriched air should be almost similar to the photosynthetic conditions when
photorespiration is depressed, i.e. low O2 and high temperature.
The present work was a study of the effect of CO2 enrichment on photo-
synthesis and photorespiration. The long-term effect of CO2 enrichment on
growth and yield was also examined. The results are discussed in relation to
the hypothesis that CO 2 enrichment is the result of a depression in photores-
piration.

MATERIAL AND METHODS

Plant cultivation. --Tomato plants (L ycopersicum esculentum cultivar 'Virosa')

were cultivated in two different ways depending on how they were to be used.
Plants used for measuring photosynthesis of attached leaves or leaf discs came
from seeds germinated in vermiculite arid transplanted to pots of rockwool
(Grodan DM 4s42/20 Hedehusene, Denmark) after 14 days. They were grown
in the p h y t o t r o n at 22/18C day/night temperature and 18 h light period at
260 pE m -2 s -1, and watered 3 times a week with 0.1% Superba nutrient solu-
tion (Nilsen, 1977).
Plants used in the growth experiments and for measurement of photosyn-
thesis on whole plants came from seeds germinated and grown in blocks of
rockwool (10 10X 6.5 cm) in an ordinary greenhouse. After germination, the
blocks were watered each day with a 0.2% Hydroflor nutrient solution (Norsk
Hydro A.S.) to which 0.05% CaNO3 had been added. After 4.5 weeks of
growth, the plants were transferred into gulleys located in one of 2 small
greenhouses and treated with ambient air or high CO2 (1000 pl CO21-1). Each
greenhouse contained 4 gulleys, 1.6 m long, with a distance of 0.4 m between
each gulley. Ten plants were placed in each gulley and 0.15% Hydroflor was
circulated through the gulley. Added to the hydroflor was: N, 95.95; P, 17.7;
K, 111.0; Ca, 98.5; Mg, 22.5; S, 34.8; Fe, 2.81; Mn, 0.05; B, 0.5; Zn, 0.06;
Cu, 0.025; Mo, 0.01 mg 1-1. The nutrient solution was changed 2--3 times a
week. Between changes, the pH was measured and corrected with KOH to a
value higher than 5. Through each gulley, 150 1 nutrient solution was circulated
at a rate of 400--600 ml min -1. The nutrient solution was maintained at 20C
and was k e p t saturated with O2 b y bubbling air through it.

Greenhouse construction. - - Two small greenhouses, 5 m 2, volume 10 m 3,

were constructed of aluminum frames covered with glass (Fig. 1). House A
(control house) had a ventilator in the ceiling and another in the door (0.25
m2); both were opened gradually by an automatic gas cylinder when the tem-
perature exceeded 18C. House B had its glass plates sealed with silicone. It
was kept closed and the CO2 concentration was maintained at 1000 + 200 pl
CO21-'. CO2 was injected from containers provided by Norsk H y d r o and
passed over an electric valve regulated by a CO2 analyzer (URAS II. Hartman
and Braun) which monitored the CO~ level.
The temperatures of Houses A and B were kept at the same level by an air-
temperature control unit. When the temperature in House B, which was kept
closed, exceeded the temperature in House A, the cooling unit started. How-
ever, the inside of the greenhouses was ultimately dependent u p o n the out-
door temperatures within the limits of the heating-capacity and ventilation
cooling of House A. The relative humidity of the houses was recorded during
the growth period and was n o t significantly different within the limit of +10%.

Fig. I. Basic greenhouse design. House A was made of glass plates and was ventilated by
openings in the ceilingand door. House B was made of glass plates which were sealed with
silicone to make the house air-tight.(V) valve for CO~ injection;(C~) unit for measuring
and regulating C O 2 level; (T A ) unit for measuring the temperature in House A; (TB)
unit for measuring the temperature in House B; (C2) unit for regulating the temperature
in House B; (A.c.u.) air conditioning unit; (S) Hartman and Braun gas switching unit.

Photosynthetic measurements. - - Measurements of photosynthesis of whole

plants or attached leaves were made in a semi-closed gas exchange system. The
automatic control and injection systems have been described by Nilsen et al.
(1975), and the gas exchange system b y Nilsen and Mortensen (1978). The
temperature was maintained at 22.5C and the relative humidity at 70%. The
assimilation chamber was illuminated from above by a 6000 W xenon lamp
(Osram XBF 6000 W/L). Different light intensities were obtained b y the use
of metal lattices and plastic screens which did n o t change the spectral distribu-
tion of the light. The light intensity was measured b y a Li-Cor Model Li-185
and a quantum sensor (Lambda Instrument Corp.).
The part of the plant above the growth substrate, or an attached leaf, was
placed in the assimilation chamber. The whole plants were 4 weeks old and
the leaves of 8-week-old plants were used. In experiments using attached leaves,
the plant was intact and was supported in such a way that leaf n u m b e r 2
could be placed in the assimilation chamber. The plants were given an adapta-
tion period o f 1 h before measurements were made. When the CO2 concentra-
tion or temperature was changed, another adaptation period of 1 h was used
before a new measurement was made, b u t when the light intensity was changed
only a 15-min adaptation period was allowed.
Net photosynthesis of 12 leaf discs, 1.7 cm in diameter, taken from the 3
lowest leaves of 5-week-old t o m a t o plants, was measured in a closed gas-exchangl
system similar to that described b y Nilsen et al. (1979). A Schlumberger (Anir
12) gas analyzer and a water-cooled assimilation chamber were used (Fig. 2).
The volume of the whole system was 0.33 1. The temperature of the assimila-
tion chamber was kept constant b y a circulating water bath. After a 1-h adap-
tation period, photosynthesis was measured.

temperature
sensor
water
gas . gas
water
water

water >

Fig. 2. Assimilation chamber used t o measure net photosynthesis of leaf discs at different
temperatures.

When net photosynthesis was measured at different temperatures and in

high CO2, the plants or plant parts were k e p t at a CO2 concentration of 1000
#1 CO21 -I between each measurement. While measuring photosynthesis, the
CO 2 level d r o p p e d to 300 ~1 CO21-1. However, the photosynthetic rate was
calculated at 480 ~l CO21-1. When the measurements were performed in low
CO2, the plants were k e p t for 1 h in air between measurements and the photo-
synthetic rate was calculated at 200 #1 CO21-1.
RESULTS AND DISCUSSION

Net photosynthesis of attached leaves of tomato plants, which had been

grown at 2 different light intensities, 130 and 260 #E m-: s-1, was measured as
CO2 uptake in a closed gas-exchange system. Figure 3 shows the rate of photo-
synthesis at different CO: concentrations and light intensities. Photosynthetic
rates of the 2 groups of plants were not significantly different at the highest
and m e d i u m level light intensities, although the light intensity was 22--34%
lower than that used for growing of plants at 130 #E m-: s-1. This indicates
that those plants grown in low light intensity had a higher photosynthetic effi-
ciency at the light intensities in which photosynthesis was measured.
Net photosynthesis increased significantly with increasing CO: concentra-
tion at all light intensities (Fig. 3). The increase in absolute values of photo-
synthesis was highest at the highest light intensity, from 10 to 16 mg CO:
dm -2 h -~, which is a 59% increase, while a 33% increase was found at 81/~E
m -2 s-~ (Fig. 3). Even at the lowest light intensity, production can be in-
creased significantly by increasing the CO: level. It is widely accepted that the
effect of CO2 enrichment is negligible at low light intensity. Gaastra (1959)
concluded that CO: enrichment only stimulates photosynthesis at high light
intensities. However, an examination of his data shows a 29% increase in pho-
tosynthesis of t o m a t o plants at 1.6 X 104 erg s-~ cm -2 when the CO2 level is
raised from 300 to 1300 pl CO: 1-1.

20

7
.:

~' 15

(..)

- 10

~ 5

00 5 0' 0 1 0 '0 0 15'00 2 0 0' 0

CO 2 c o n c e n t r a t i o n , IJl-1-1

Fig. 3. Net p h o t o s y n t h e s i s by attached leaves of t o m a t o plants measured in different light

intensities: 96 (o), 193 (a), 81 (o), 130 ( . ) , and 236 ( - ) / ~ E m -~ s-'. The plants were grown
in 140 ~E m -~ s -~ (open symbols) or 260/~E m "~ s-' (closed symbols) in climate-chambers
illuminated with Philips T L M F 33 RS 140W for 2 m o n t h s b e f o r e m e a s u r e m e n t s were made.
Each point represents 2--3 measurements.
 It seems apparent that CO2 enrichment of plants also reduces photorespira-
tion. The increased photosynthesis and production in response to CO2 enrich-
ment observed in the present study suggests that photorespiration is reduced.
The present view is that the CO2/O2 ratio determines the balance between
photosynthesis and photorespimtion (Bowes et al., 1971). CO: and O2 work
as competitive molecules on the enzyme site of RuBP-carboxylase/oxygenase
(Bowes and Ogren, 1972; Lorimer et al., 1973). Both the oxygenase reaction
(photorespiration) and the carboxylase reaction (photosynthesis) are part of
the carbon reduction cycle and require energy (ATP--NADPH) from the light
reaction. Hicklenton and Jolliffe (1978) found that CO2 enrichment produces
a lower CO2 compensation point, which is a further indication of a reduction
in photorespiration. A reduction in glycolic acid oxidase was observed in to-
mato plants treated with high CO2, and long-term treatment with high CO2 in-
creases the activity of RuBP-carboxylase (Fair et al., 1973).

Net photosynthesis of whole tomato plants was measured in 21% 02 (air)

and in 2% 02 at 2 different CO2 concentrations. Photosynthesis increased
when the measurements were performed in an atmosphere with a high CO2/O2
ratio (2% O2 or 1000 ul CO21-1) where the oxygenase reaction was depressed
(Table I). However, these same conditions did not increase photosynthesis sig-
nificantly. This indicates that a CO2 concentration of 1000 ~1 CO21-1 stimu-
lates photosynthesis by depressing photorespiration to a similar extent as in
2% O2, which further strengthens the view that increasing the CO2 concentra-
tion alters the competitive inhibition of the RuBP carboxylase enzyme in
favour of photosynthesis. Table I also shows that the stimulation in photosyn-
thesis obtained by reducing 02 or increasing the CO2 concentration occurred
in both light intensities. Therefore, a practical application of these data would
mean that CO2 enrichment should be given during the entire light period in
order to obtain maximum production. This is consistent with data presented
by Calvert and Slack (1976), who showed that CO2 given to plants at sunrise,

TABLE I
The effect of high and low CO2and O~ concentrations and light i n t e n s i t y o n net p h o t o s y n -
thesis (rag CO 2 g-~ h -~) o f w h o l e plants cultivated in a m b i e n t CO2 (air)

Light 02 #1 CO2 1-~ Increase

intensity (%) (%)
330 2700
350~Em ~ s -1 21 17 5 29 8 67
2 28 3 33 7 17
Increase (%) 63 15

585~Em ~ s -1 21 26 5 40 9 53
2 39 7 4 4 11 11
Increase (%) 50 9
when the light intensity is very low b u t above the light compensation point,
significantly increased the yield as compared with CO2 enrichment provided
later in the day.

The effect o f long-term treatment in C02 enriched air on photosynthetic

rate was measured at 2 CO2 levels (330 and 1000 pl CO21 -t) on t o m a t o plants
grown in air or in CO2-enriched air. Plants cultivated in air showed a higher
net photosynthesis when measured in 1000/~l CO21-1 than plants cultivated at
this high CO2 level (Fig. 4). Net photosynthesis was lower for plants cultivated
in high CO2 than for plants cultivated in air when the measurements were
made in 330 pl CO21-1. Similar results were observed by Ho (1977) on fully
developed t o m a t o leaves (No. 7), while Madsen (1971) found that photosyn-
thesis b y 1-month-old t o m a t o plants cultivated in high CO2 (1000--5000 pl
CO21 -t) was higher when measured at 300--500/A CO21 -t. Rice plants culti-
vated in high CO2 showed a reduced net photosynthesis in 300 pl CO21 -t
(Imai and Murata, 1978). Results similar to those from the present investiga-
tion were reported for radish b y Frydrych (1976).

~. 100

-o
o~ 80

~ 6o

}~-,o~oo~

o 20

Z I i i i i

0 100 200 300 400 500

Light intensity, ~E m-2s -1

Fig. 4. Net photosynthesis of whole plants at different light intensities and 2 002 concen-
trations: 3 102 (aA) and 103 (oo) ~1 CO 21-~. The plants had been grown for 1 month in
either ambient CO 2 (open symbols) or 103 ~1 CO 21-~ (closed symbols). The data represent
the averages of 3 measurements.

The decrease in photosynthesis when plants are grown in high CO2 and
when photosynthesis is measured in atmospheric CO2 has been attributed to
an accumulation o f photosynthetic products or starch. Madsen (1968) ob-
served an increased a m o u n t of starch in t o m a t o leaves with increasing CO2
concentration. Ito (1971) showed that an increased starch content reduced
net photosynthesis, and he suggested that this was caused by an increase in
mesophyll resistance.
Normally, the temperature of greenhouses cannot be controlled. The tem-
perature can easily be k e p t constant at the lower temperature-range by
heating, b u t on sunny days the heat radiation exceeds the cooling effect of
ventilation and temperatures of 30C around the plants and 60C at the ceiling
are common. Even in February, the greenhouse temperature in Norway can
exceed 25C, and on sunny days there may be a need for ventilation when the
outdoor temperature is several degrees below zero. Therefore, plants are often
cultivated in temperatures above the optimum.

Net photosynthesis of leaf discs was measured at different temperatures

and in 2 different CO: concentrations (330 and 1000 pl CO21-1). The plants
had been cultivated in atmospheric CO2 previous to the measurements. The re-
sults (Fig. 5) indicate a broad optimal temperature between 15 and 20C for
net photosynthesis in 330 ~1 CO21-1. At the higher CO2 level of 1000 pl CO2
1-1, the optimal temperature was a little higher, about 25C. In this case, the
slope of the photosynthetic curve is much steeper on each side of the optimal
temperature, which indicates that there is a sharp reduction in photosynthesis
at non-optimal temperatures. Similar results were obtained by Bj~brkman et al.
(1978) on Nerium oleander when net photosynthesis was measured under
CO2-saturating conditions.
The increase in optimal temperature for net photosynthesis in CO2-enriched
air may be attributed to the inhibiting effect of CO2 on photorespiration. The
temperature optimum of net photosynthesis in air (21% 02 and 350 ~l CO:
20

18

~= 14

o~" 12,
o /t/]} 1000 NI CO2

z 4
350 ~11CO

0 5 10 15 20 25 30 35 40 45
Temperature, C

Fig. 5. Net photosynthesis of leaf discs after adaptation to different temperatures and 2
different CO 2 concentrations, 350 and 1000 ~1 CO 214. The actual photosynthetic rates
were calculated at 200 and 480 ~1 CO 214, respectively.
1-1 ) is, in reality, a combination of the optimal temperature of the oxygenase
and carboxylase reactions of RuBP. When the oxygenase reaction is inhibited
by high CO2 or low O:, the optimal temperature for net photosynthesis moves
upwards, since the RuBP oxygenase reaction has a higher optimal temperature
than the RuBP carboxylase reaction (Laing et al., 1974; Badger and Andrews,
1974; Peisker and Apel, 1977). Another way to inhibit photorespiration is by
decreasing the O2 concentrations (BjSrkman, 1966), for example a decrease in
the O2 level from 31 to 3% increased the temperature optimum for net photo-
synthesis of Triticum sativum from 20--26C to 30--36C (Jolliffe and
Tregunna, 1968). Photosynthesis in C4 plants, in which photorespiration is
either lacking or depressed, has a temperature optimum that is higher than Ca
plants (Long et al., 1975). However, the temperature response of photosyn-
thesis at various concentrations of O: and CO: can only partially be determined
from the kinetic properties of RuBP carboxylase/oxygenase reactions. Berry
and Bj6rkman (1980) point out that the temperature response of the photo-
synthetic electron transport system and the relative capacity of the carboxyla-
tion reaction to receive electrons must also be considered. The practical appli-
cation of these findings is that plants can be cultivated optimally in a higher
temperature when they are grown in CO2-enriched air than when grown in air.

Plant production in greenhouses enriched with CO:. -- Short-term photosyn-

thetic measurements are useful when studying mechanisms of growth and the
effect of different climate parameters, but the information obtained cannot
always be converted into practical applications. An experiment with a practi-
cal application was the study of plant growth and yield when enrichment was
provided during the entire growth period, which was compared with growth
under similar climatic conditions in atmospheric CO2.
Fresh weight, dry weight and percent dry matter of 10 plants after a culti-
vation period of 30 or 99 days were determined. The leaves and stems were re-
moved and weighed separately. After 30 days growth, CO2 enrichment had a
significant effect on fresh weight, dry weight and percent dry matter of the
whole plant, leaves and stems (Table II). The most significant finding was that
dry weight, and thus percent dry matter, were increased the most, which is a
clear indication that there was a real increase in production.
The lack of a significant difference in production between the control plants
and CO2-enriched plants after 99 days (Table II) might be due to a saturation
in the level of production in high CO2 followed by a decline in growth rate.
The plants cultivated in normal CO2 may have kept a lower, but a more con-
stant, growth rate, but after 99 days the same level is reached as the CO2-en-
riched plants. Hurd (1968) observed a similar tendency. The relative growth
rate of young tomato plants treated with high CO2 was higher, but the leaf-
area-ratio decreased, which is an indication that the excess photosynthetic
products were not used for the production of new leaves, but were placed in a
pool which could have been used for the development of inflorescences and
fruits. Therefore, the increase in photosynthesis by CO: enrichment would
accelerate these processes.
10

TABLE II

The effect of CO 2 enrichment (1000 ~1 CO 2 1-1) on the growth of tomato plants in House
B. Fresh weight (Fw), dry weight (Dw) and percent dry matter (D%) of the treated plants
were calculated as percent of control plants. Both groups of plants were grown at the same
temperature. Ten plants were harvested at each growth period. The difference was statisti-
cally significant at 99% for all growth measurements after 30 days, but no significant dif-
ference was found after 99 days growth.

30-day growth period 99-day growth period

Fw Dw D% Fw Dw D%

Whole plant 30 60 21 0.3 10 9

Leaves 27 57 25 2.4 8 6
Stems 36 67 22 -4.2 13 16

CO 2 treatment does not seem to change the weight distribution of the dif-
ferent parts of the plant (Table II). This is also consistent with the findings of
Hurd (1968).
The tomatoes were harvested as they became ripe, counted and weighed.
After a harvest period of 1 month, all remaining tomatoes were harvested, sep-
arated into ripe and unripe, and the total yield was determined (Table III).
The effect of CO2 enrichment was most pronounced on the yield of ripe to-
matoes; 89% increase. The total yield was increased by 29%.
It is difficult to compare growth experiments by different investigators, be-
cause in most cases the experimental conditions are quite different. Calvert
(1972) found a 90% increase in yield after 4 weeks growth of tomato in high
CO2, but these plants were treated with high CO2 even after the pricking-out
stage. When plants were used which had been transferred at the same develop-
mental stage and treated with CO2 in a manner similar to that used in the
present investigation, an 80% increase in yield after 4 weeks was obtained and
only a 37% increase after 20 weeks. CO2 enrichment was discontinued after
the 4-weeks harvesting, and the reduction in yield at the end of 20 weeks may
have been due to the reduction in CO2 concentration. Since the plants were
cultivated during winter, light could also have been a limiting factor.
Hicklenton and JoUiffe (1978) provided CO2 enrichment to tomato plants
during the entire daylight period, but ventilation became necessary when the
room temperature exceeded 24C and CO2 enrichment had to be discontinued.
CO2 enrichment was begun as soon as seedlings emerged and was continued
until late April, which means that no CO2 was given after the first fruits were
harvested in mid-May.An increase in yield of 45% can be calculated from
their data in response to high CO2 (900 #l CO21-1) for 4 weeks. This yield is
low in comparison to the yield obtained in the present investigation (Table III).
We added CO2 not only during the entire light period, but also during the en-
tire growth period, including flower and fruit development. Recently, Clough
and Peet (1981) also found that continuous enrichment with CO2 is more
 11

beneficial than intermittent treatment. Therefore, it appears that greater yield

can be obtained when the greenhouse is kept closed and CO2 is added continu-
ously during the day.
The 23% increase in the number of tomatoes per plant was significant (Table
III). Morgan (1971) reported a 15% increase and Calvert and Slack (1975) also
observed a significant increase in number of tomatoes by plants treated with
high CO2. The increase in the weight of each tomato was small and not signifi-
cantly different from the control group. This finding is consistent with that
reported by Wittwer and Robb (1964), who found an increase in fresh weight
per tomato of 15% when plants were cultivated in high CO: (800--2000 ~1
CO 21-1). However, Kimball and Mitchell (1979) found no change in the size
of tomatoes cultivated in 1000 pl CO21-1.

TABLE III

Yield o f t o m a t o e s in t w o g r e e n h o u s e s : C o n t r o l H o u s e A w i t h air v e n t i l a t i o n ; H o u s e B en-

r i c h e d w i t h CO2 ( 1 0 0 0 ~1 CO21-1). H o u s e B was k e p t closed a n d t h e t e m p e r a t u r e m a i n -
t a i n e d a t t h e s a m e level as t h a t in t h e c o n t r o l h o u s e b y a c o o l i n g / h e a t i n g system. D a t a
are t h e m e a n SD o f 9 c o n t r o l p l a n t s a n d 8 p l a n t s t r e a t e d w i t h h i g h CO 2 . * S i g n i f i c a n t l y
d i f f e r e n t f r o m H o u s e A, > 95%

House A House B
control (CO 2 enriched)

F r e s h w e i g h t t o m a t o e s p e r p l a n t (kg)
Ripe 1.4 0.7 2.6 0.6*
Unripe 1.6 + 0.5 1.3 0.4
Total 3.0 0.9 3.9 +- 0.7*

Number of tomatoes per plant

Ripe 17 9 34 8*
Unripe 40 9 36-+ 7
Total 57 15 70 13

F r e s h w e i g h t p e r t o m a t o (g)
Ripe 79 27 76 + 32
Unripe 41 27 35 + 35
Total 52 32 55 -+ 34

The total yield (gram per plant or number of tomatoes per plant) showed a
significantly positive effect in response to CO2 enrichment provided through-
out the entire harvest period (Fig. 6). Hicklenton and Jolliffe (1978) observed
a similar trend, but the effect was n o t so pronounced, especially during the
first part of the harvest period. This may have been caused by the fact that
the plants were given high CO2 only up to 14 days before the harvest period.
The quality of the tomatoes was n o t significantly different in the 2 CO:
concentrations (data not shown). Calvert and Slack (1975) found a small,
though significant increase in the percentage of select grade fruits when CO2
enrichment was given during the pre-planting-period, but no difference when
CO2 enrichment was provided only during the post-planting-period.
12

103 rUl CO 2 ]

? 2.0 /
v~
/
:" 1.0
.~- /" o/

05
/e--e
0 ~- T I i I I I i i i i
6 9 12 15 18 21 24 27 30 2
Aug. I Sept.

Fig. 6. The cumulative yield of tomato plants in response to CO~ enrichment, The data rep,
resent the mean kg weight of fruits per plant. The control house was ventilated with air.
The CO s house was kept air-tight and enriched with 10 3ul COs 1-~ during the entire growing-
period. The temperature of the COs house was kept at the same level as that in the control
house by a cooling/heating system.

ACKNOWLEDGEMENTS

We gratefully acknowledge the valuable assistance of Dr. M.K. Haugstad in

the writing of this manuscript, and the preparation of the figures by L.A.
Oritsland. This work was performed as a part of the project: "CO2 and heat
from charcoal in greenhouses", initiated by the Norwegian Defence Research
Establishment, and was supported by a grant from the National Agricultural
Research Council and the Royal Ministry of Petroleum and Energy of
Norway.

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